Acta zoologica cracoviensia, 58(2): 137-171, Kraków, 31 December, 2015 Ó Institute of Systematics and Evolution of Animals, Pol. Acad. Sci., Kraków doi:10.3409/azc.58_2.137
Recapitulation of data on Ukrainian fossil insectivore mammals (Eulipotyphla, Insectivora, Mammalia)
Barbara RZEBIK-KOWALSKA and Leonid I. REKOVETS
Received: 04 September 2015. Accepted: 08 December 2015. Available online: 23 December 2015.
RZEBIK-KOWALSKA B., REKOVETS L.I. 2015. Recapitulation of data on Ukrainian fossil insectivore mammals (Eulipotyphla, Insectivora, Mammalia). Acta zool. cracov., 58(2): 137-171. Abstract. The paper recapitulates data on the fossil insectivore mammals of Ukraine. They include families: Erinaceidae, Talpidae, Dimylidae, Heterosoricidae and Soricidae. In Ukraine the oldest insectivore remains come from the locality of Gritsev and are dated to the early Late Miocene (MN9). The insectivore taxa cited from this country are presented in twelve Tables of three types. One of them lists taxa published with descriptions, meas- urements, localities and geological ages, the second mentions taxa without descriptions or measurements and the third without descriptions, measurements or exact localities. Short remarks on questionable forms are given. Key words: Erinaceidae, Talpidae, Dimylidae, Heterosoricidae, Soricidae, Late Miocene-Holocene, Ukraine. * Barbara RZEBIK-KOWALSKA, Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, S³awkowska 17, 31-016 Kraków, Poland. [email protected] Leonid I. REKOVETS, University of Environmental and Life Sciences of Wroc³aw, Che³moñskiego 38 C, 51-630 Wroc³aw, Poland; National Museum of Natural History, B. Khmelnitski 15, 01030 Kiev, Ukraine. [email protected]
I. INTRODUCTION
Many papers were published on fossil mammals of Ukraine but only a dozen or so con- tain description of insectivore mammals with measurements, systematic positions as well as localities in which they were found and their geological ages. Those are the papers of PIDOPLICHKO 1955, 1956, TATARINOV 1958, TOPACHEVSKY 1959, 1962a, 1965 and 1966, SVISTUN 1968, MEZHZHERIN and SVISTUN 1968, GUREEV 1971, 1979, MEZHZHERIN 1972, TOPACHEVSKY and PASHKOV 1983, 1990, PASHKOV and TOPACHEVSKY 1990, RZEBIK-KOWALSKA and TOPACHEVSKY 1997, AGADZHANYAN 2009, RZEBIK-KOWAL- SKA and NESIN 2010 and RZEBIK-KOWALSKA and REKOVETS in press. Other papers con- cerning Ukrainian localities, their stratigraphy and fauna cite only the lists of insectivore taxa. The lists are usually copied from other publications of the same type, however with- out any verification. On the other hand it is known that some of these taxa were wrongly identified or improbable in Ukrainian territory. B. RZEBIK-KOWALSKA, L. I. REKOVETS 138 As it can be seen in the Tables (I, IV, VIII, X) so far 40 named species are cited but only 25 (ten moles and 15 shrews) of 15 genera (four moles and eleven shrews) are well docu- mented. As in neighbouring and territorially smaller Poland the number of well docu- mented fossil insectivore mammals (Erinaceidae, Talpidae, Dimylidae, Heterosoricidae and Soricidae) equals 77 (RZEBIK-KOWALSKA 2009) the Ukrainian list seems to be in- complete. The present paper lists in Tables (I-XII) all fossil insectivore mammals described or cited at any time in this country. It also shows which of them are well documented and really present on this area. Short remarks on the questionable forms are also given. Abbreviations: LM = Late Miocene, P = Pliocene, EP = Early Pliocene, LP = Late Pliocene, Pl = Pleistocene, EPl = Early Pleistocene, MPl = Middle Pleistocene, LPl = Late Pleistocene, H = Holocene and EH = Early Holocene. Surnames and initials of three authors are spelled in tables and references following original publications in Russian or Ukrainian, respectively: CHEPALYGA (in Russian pa- pers), CHEPELIGA (in Ukrainian); V. A. MEZHZHERIN (in Russian), V. O. MEZHZHERIN (in Ukrainian); V. A. TOPACHEVSKY (in Russian) and V. O. TOPACHEVSKY (in Ukrainian).
II. FOSSIL INSECTIVORE MAMMALS OF UKRAINE
Class: Mammalia LINNAEUS, 1758
Superorder: Insectivora sensu NOVACEK, 1986
Order: Eulipotyphla WADDELL, OKADA and HASEGAWA, 1999
II.1. Family Erinaceidae FISCHER, 1814 (Tables I-III) Comments. In the Ukraine the fossil Erinaceidae are represented by two subfamilies – the spineless gymnures – Galericinae and spiny hedgehogs – Erinaceinae. The presence of the fossil hedgehog Lanthanotherium FILHOL, 1888 is not confirmed by morphological studies. Because it is found in many European countries situated in the vicinity of Ukraine, e.g. in Hungary, Slovakia and Romania, its presence in Ukraine is also very probable. Its occurrence in Europe was documented since the Middle to the middle Late Miocene (MN3/MN4 to MN11) (RZEBIK-KOWALSKA 2009) and in Asia (China) from MN4 and MN11-MN12 (QIU and STORCH 2005). The second fossil form, Amphechinus AYMARD, 1850 from Gritsev also lacks a mor- phological description. As the remains of this genus were collected in western and south- ern Europe in localities dated from the Late Oligocene to the Middle Miocene (MN9), its presence in Ukraine is possible. It is also known from North America, Africa and Asia (ZIEGLER 2005). The name of the tribe, Amphechini GUREEV, 1979, cited by TOPACHEVSKY et al. (1997) is used very rarely in the European literature. Recapitulation of Ukrainian fossil insectivores 139
The recent Hemiechinus auritus (GMELIN, 1770) currently lives in the steppe zone of eastern Ukraine and its remains among the Late Pleistocene and Holocene fossils of this country are expected. Instead, the presence of Erinaceus europaeus LINNAEUS, 1758 cited by TOPACHEVSKY (1957a, 1961b) in the Middle and Late Pleistocene in Ukrainian territory is unlikely. This European hedgehog is distributed in western Europe and the eastern boundary of its range reaches only the Odra river. On the other hand in the Pleistocene several other species of the genus Erinaceus occurred in Europe. Moreover, the white-breasted hedgehog, E. rou- manicus BARRET-HAMMILTON, 1900 currently inhabits eastern Europe and the Ponto- Mediterranean region (SOMMER 2007). The remains mentioned by TOPACHEVSKY (1957a, 1961b) probably belong to the latter species. Erinaceidae gen. et sp. indet. from Popovo 3 and Verkhnya Krynitsa 2 are now de- scribed as Schizogalerix sp. Hedgehog remains were not found in the material from Popovo 2.
Table I Fossil Erinaceidae of Ukraine (published with descriptions, measurements, localities and geological ages)
Family Subfamilies Localities Age Publications Genera Species
Erinaceidae
Galericinae
Schizogalerix cf. sarmaticum Mikhailovka 1 LM (Late Vallesian, MN10) RZEBIK-KOWALSKA and UNGU ESIN (L , 1981) Frunzovka 2 LM (Early Turolian, MN11) N 2010
Popovo 3 LM (Early Turolian, MN11) RZEBIK-KOWALSKA and Schizogalerix sp. LM (Early/Middle Turolian, EKOVETS Verkhnya Krynitsa 2 R in press MN11/MN12)
Erinaceinae
RZEBIK-KOWALSKA and cf. Postpalerinaceus sp. Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2010
Erinaceus sp. Nogaisk EPl (Early Biharian, Q1) V. A. TOPACHEVSKY 1965 B. RZEBIK-KOWALSKA, L. I. REKOVETS 140 Table II Fossil Erinaceidae of Ukraine (mentioned without descriptions or measurements)
Family Subfamilies Tribes Localities Age Publications Genera Species
Erinaceidae
Galericinae
V. A. TOPACHEVSKY, NESIN and I. V. TOPACHEVSKY 1997; RZEBIK-KOWALSKA and V. O. Lanthanotherium sp. Gritsev LM (Early Vallesian, MN9) TOPACHEVSKY 1997; NESIN and NADACHOWSKI 2001; NESIN 2013
Schizogalerix cf. Mikhailovka 1 LM (Late Vallesian, MN10) NESIN 2013 sarmaticum Frunzovka 2 LM (Early Turolian, MN11)
V. A. TOPACHEVSKY, NESIN and I. V. TOPACHEVSKY 1997; LM (Early Vallesian, Gritsev RZEBIK-KOWALSKA and V. O. MN9) TOPACHEVSKY 1997; NESIN Schizogalerix sp. and NADACHOWSKI 2001
Novoelizavetovka 3 LM (Middle Turolian, MN12) NESIN 2013
Cherevychnoe 3 LM (Middle Turolian, MN12) NESIN 2013
V. A. TOPACHEVSKY, NESIN Galericinae gen. and I. V. TOPACHEVSKY 1997; Gritsev LM (Early Vallesian, MN9) et sp. indet NESIN and NADACHOWSKI 2001
Erinaceinae
V. A. TOPACHEVSKY, NESIN ?Amphechini Gritsev LM (Early Vallesian, MN9) and I. V. TOPACHEVSKY 1997
RZEBIK-KOWALSKA and V. O. Gritsev LM (Early Vallesian, MN9) TOPACHEVSKY 1997; NESIN Amphechinus sp. 2013
Cherevychnoe 3 LM (Middle Turolian, MN12) NESIN 2013 cf. Hemiechinus sp. Gorishnya Vygnanka LP TATARINOV 1965b cf. Postpalerinaceus sp. Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 Recapitulation Ukrainian fossil insectivores 141 Table II (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species
Kotlovina 1 EP (Late Ruscinian, MN15)
Odessa (catacombs) EP (Late Ruscinian, MN15)
EPl (previously Late ESIN Kotlovina 2 N 2013 Villanyan, MN17)
EPl (previously Late Kotlovina 3 Villanyan, MN17)
Nogaisk EPl (Early Biharian, Q1) V. O. TOPACHEVSKY 1957b
Erinaceus sp. REKOVETS, CHEPALYGA, Bolshevik 2 (l. III) EPl (Late Biharian, Q2) NESIN and SVETLITSKAYA 1990; REKOVETS 1990
Tarkhankut EPl (Early Biharian, Q1)
Nogaisk EPl (Early Biharian, Q1) REKOVETS and Cherevychnoe 1 EPl (Early Biharian, Q1) NADACHOWSKI 1995 Bolshevik 2 (l. III) EPl (Late Biharian, Q2)
Tikhonovka 1 EPl (Late Biharian, Q2)
Novoelizavetovka 2 LM (Early Turolian, MN11)
Popovo 3 LM (Early Turolian, MN11) NESIN 2013 Erinaceidae gen. LM (Early/Middle Verkhnya Krynitsa 2 et sp. indet. Turolian, MN11/MN12) REKOVETS and PASHKOV 2009
LP (Early Villanyan, end of Popovo 2 MN16) B. RZEBIK-KOWALSKA, L. I. REKOVETS 142 Table III
Fossil Erinaceidae of Ukraine (mentioned without descriptions, measurements or exact localities)
Family Subfamily Localities Age Publications Genera Species Erinaceidae Erinaceinae alluvia (lower course of LPl and H V. O. TOPACHEVSKY 1957a Dnieper) alluvia (middle course of Erinaceus europaeus Dnieper, MPl and LPl V. O. TOPACHEVSKY 1961b LINNAEUS, 1758 Cherkasy-Mishurin Rig and Kiev-Cherkasy) Podolia and Prikarpatie HTATARINOV 1970 (north to Carpathian Mts Erinaceus europaeus rumanicus (should be E. e. roumanicus) Podolia and Prikarpatie HTATARINOV 1970 now (north to Carpathian Mts) E. roumanicus BARRET-HAMMILTON, 1900 Podolia and Prikarpatie Erinaceus sp. HTATARINOV 1970 (north to Carpathian Mts) Hemiechinus auritus (GMELIN, alluvia (lower course of LPl and H V. O. TOPACHEVSKY 1957a 1770) Dnieper) Podolia and Prikarpatie cf. Hemiechinus sp. PTATARINOV 1970 (north to Carpathian Mts)
II.2. Family Talpidae FISCHER, 1814 (Tables IV-VI) Comments. Identification of Desmaninae species is difficult especially in the case of isolated teeth. Three genera of water-moles (Desmaninae) are listed for Ukraine, including Mygalinia, Ruemkelia and Desmana. The presence of the small forms Mygalinia and Ruemkelia is well documented. However, their identification [Mygalinia (=Ruemkelia)] by V.O. TOPA- CHEVSKY et al. (1997) and NESIN and NADACHOWSKI (2001) is incorrect because these two genera distinctly differ and both were found in Ukraine. Also an identification Recapitulation Ukrainian fossil insectivores 143
Mygalinia (=Dibolia) sp. (NESIN 2013) is incorrect because a new name for Dibolia RÜMKE, 1985 is Ruemkelia RZEBIK-KOWALSKA and PAW£OWSKI, 1994. The genus Ruemkelia (a new name for Dibolia RÜMKE, 1985, an invalid homonym, see RZEBIK-KOWALSKA and PAW£OWSKI 1994) is considered by some authors, e.g. HUT- TERER (1995), as a synonym of the genus Archaeodesmana TOPACHEVSKY and PASH- KOV, 1983. However, the Archaeodesmana has as its type species Desmana pontica SCHREUDER, 1940, and the genus Dibolia (=Ruemkelia) has as its type species Dibolia dekkersi RÜMKE, 1985. According to the first author of the present paper such an assump- tion (congenerity of D. pontica and D. dekkersi) should not be established without a study of original material. The new name Ruemkelia seems to be more correct because the re- placement of the name Dibolia by Ruemkelia had a purely nomenclatorical character and does not depend on the “state of art” in systematic of the desmans. So far, besides the recent Desmana moschata, ten named species of Desmana were de- scribed or identified from Ukrainian territory (see above). Five of them (D. thermalis, D. kujalnikensis, D. meridionalis, D. moldavica and D. moschata) were listed from the Late Pliocene. However, from the ecological point of view the presence of so many forms adapted to one (aquatic) environment seems improbable. The species described by TOPACHEVSKY and PASHKOV (1990) and PASHKOV and TOPACHEVSKY (1990) were not compared with known forms from Europe (e.g. D. moldavica was compared only with D. jalpugensis and D. kujalnikensis only with D. nehringi). The sizes of the desmans described by them do not differ substantially. Their holotypes were described on the grounds of different teeth (e.g. D. jalpugensis based on a mandible with p4, D. kuljakensis based on an isolated upper P3) and particular teeth and their characters were described incidentally (e.g. in p4 of D. jalpugensis the described talonid is very well developed, in D. moldavica it is poorly developed but in D. kujalnikensis this structure is not mentioned) and therefore many Ukrainian remains of this genus are still unidentified. The species Desmana pontica described by SCHREUDER (1940) from the Late Miocene Polgárdi in Hungary and cited by NESIN (2013) from Cherevychnoe 3 was transferred by RÜMKE (1985) to the genus Dibolia and now it represents Ruemkelia species (RZE- BIK-KOWALSKA and PAW£OWSKI, 1994) or Archaeodesmana (HUTTERER 1995). The presence of the genus Galemys is problematical in Ukraine. One species, Desmana kormosi, cited by REKOVETS and PASHKOV (2009) from Popovo 2 and acknowledged by RÜMKE (1985) as belonging to the genus Galemys (Galemys kormosi) may suggest its presence in Ukraine. However, in the locality Popovo 2 there were no remains belonging to the genus Galemys. It is, however, probable that some specimens identified as Desmana sp. actually represent the genus Galemys. The genus is known from neighboring Poland (MN14/MN15-MN16, RZEBIK-KOWALSKA 2009) and Hungary (Early Pleistocene, JÁNOSSY 1986). A revision of the Ukrainian desmans is necessary. The presence of the American (Late Miocene – Early Pliocene) genus Domninoides GREEN, 1956 in Europe was criticized by ZIEGLER (1999). Described by GIBERT (1975) from the Middle Miocene (MN6) of Spain, it was also noted from Gritsev by TOPACHEVSKY (in RZEBIK-KOWALSKA and TOPACHEVSKY 1997) and in the paper of NESIN and NADACHOWSKI (2001), the authors listed it as the dominant taxon. However, these remains were never studied in detail. B. RZEBIK-KOWALSKA, L. I. REKOVETS 144 Table IV Fossil Talpidae of Ukraine (published with descriptions, measurements, localities and geological ages) Family Subfamilies Tribes Localities Age Publications Genera Species Talpidae Desmaninae cf. Mygalinia hungarica LM (Early/Middle RZEBIK-KOWALSKA and Verkhnya Krynitsa 2 (KORMOS, 1913) Turolian, MN11/MN12) REKOVETS in press Mikhailovka 1 LM (Late Vallesian, MN10) RZEBIK-KOWALSKA and Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2010 Ruemkelia sp. Frunzovka 2 LM (Early Turolian, MN11) LM (Early/Middle RZEBIK-KOWALSKA and Verkhnya Krynitsa 2 Turolian, MN11/MN12) REKOVETS in press
Desmana aff. nehringi Novopetrovka EP (Early Ruscinian, MN14) V. O. TOPACHEVSKY KORMOS, 1913 Velikomikhailovka EP (Early Ruscinian, MN14) 1962a Odessa, 16th Station of RZEBIK-KOWALSKA and Desmana cf. nehringi LM (Late Turolian, MN13) Bolshoy Fontan NESIN 2010 V. O. TOPACHEVSKY Nogaisk EPl (Early Biharian, Q1) Desmana thermalis 1962a, 1965 KORMOS, 1930 V. O. TOPACHEVSKY Kairy EPl (Early Biharian, Q1) 1962a Desmana (Pliodesmana) EPl (previously Late PASHKOV and V. A. jalpugensis PASHKOV and Kotlovina (probably 2, 3) Villanyan, MN17) TOPACHEVSKY 1990 TOPACHEVSKY, 1990 LP (Early Villanyan, end Desmana Cherevychnoe 2 (Galemodesmana) of MN16) PASHKOV and V. A. kujalnikensis PASHKOV LP (Early Villanyan, TOPACHEVSKY 1990 Kryzhanovka 2 and TOPACHEVSKY, 1990 middle of MN16)
Desmana meridionalis Zhevakhova Gora (ls. LP (Early Villanyan, V. A. TOPACHEVSKY and TOPACHEVSKY and 11, 15) middle of MN16) PASHKOV 1990 PASHKOV, 1990 Tiligul (=Morskoe) EPl (Early Biharian, Q1) Cherevychnoe 1 EPl (Early Biharian, Q1) Desmana nogaica V. A. TOPACHEVSKY and TOPACHEVSKY and Nogaisk EPl (Early Biharian, Q1) PASHKOV 1990 PASHKOV, 1990 Taman EPl (Q1) Recapitulation Ukrainian fossil insectivores 145 Table IV (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Desmana gureevi Kara0 Dubina EPl (Early Biharian, Q1) V. A. TOPACHEVSKY and TOPACHEVSKY and PASHKOV 1990 PASHKOV, 1990 Luzanovka EPl (Early Biharian, Q1) PIDOPLICHKO 1956 Desmana moschata Sinyakovo EPl ternopolitava V. A. TOPACHEVSKY 1962a PIDOPLICHKO, 1956 Gorishnya Vygnanka LP TATARINOV 1958 V. O. TOPACHEVSKY Desmana moschata Novogrod-Severskii LPl (Late Weichselian) palaeoborysthenica 1959, 1962a TOPACHEVSKY, 1959 Romankovo MPl SVISTUN 1968 LP (Early Villanyan, Verkhnya Krynitsa 1 beginning of MN16) LP (Early Villanyan, end Popovo 2 RZEBIK-KOWALSKA and Desmana sp. of MN16) REKOVETS in press P/Pl boundary (previously Popovo 1 Early/Late Villanyan, MN16/MN17) LM (Early/Middle RZEBIK-KOWALSKA and cf. Desmana sp. Verkhnya Krynitsa 2 Turolian, MN11/MN12) REKOVETS in press Desmaninae gen. et sp. Frunzovka 2 LM (Early Turolian, MN11) indet. 1 RZEBIK-KOWALSKA and Desmaninae gen. et sp. NESIN 2010 Mikhailovka 1 LM (Late Vallesian, MN10) indet. 2 Uropsilinae Desmanella cf. dubia RZEBIK-KOWALSKA and Mikhailovka 1 LM (Late Vallesian, MN10) RÜMKE, 1976 NESIN 2010 RZEBIK-KOWALSKA and Desmanella sp. Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2010 Talpinae Urotrichini Urotrichus giganteus RZEBIK-KOWALSKA and Mikhailovka 2 LM (Early Turolian, MN11) ZIEGLER, 2006 NESIN 2010 RZEBIK-KOWALSKA and ?Scaptonyx sp. Mikhailovka 1 LM (Late Vallesian, MN10) NESIN 2010 B. RZEBIK-KOWALSKA, L. I. REKOVETS 146 Table IV (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Talpini Talpa gilothi STORCH, RZEBIK-KOWALSKA and Mikhailovka 2 LM (Early Turolian, MN11) 1978 NESIN 2010 Talpa cf. europaea Srednyaya Cave Pl TATARINOV 1965a LINNAEUS, 1758 (near Nizhnee Krivche) Talpidae incertae sedis Desmanodon major RZEBIK-KOWALSKA and Mikhailovka 2 LM (Early Turolian, MN11) ENGESSER, 1980 NESIN 2010
Table V Fossil Talpidae of Ukraine (mentioned without descriptions or measurements) Family Subfamilies Tribes Localities Age Publications Genera Species Talpidae Desmaninae V. A. TOPACHEVSKY, Migalinia sp. Mikhailovka 2 LM (Early Turolian, MN11) NESIN and I. V. TOPA- (should be Mygalinia sp.) CHEVSKY 1997 NESIN and Mygalinia sp. Mikhailovka 2 LM (Early Turolian, MN11) NADACHOWSKI 2001 V. A. TOPACHEVSKY, LM (Late Vallesian, NESIN and I. V. TOPA- Mikhailovka 1 Mygalinia (=Ruemkelia) MN10) CHEVSKY 1997; NESIN and NADACHOWSKI 2001 Vinogradovka 1 LM (Late Turolian, MN13) NESIN 2013 Ruemkelia (=Mygalinia) Mikhailovka 1 LM (Late Vallesian, MN10) NESIN 2013 Mygalinia (=Dibolia) sp. Novoelizavetovka 2 LM (Early Turolian, MN11) NESIN 2013 Mikhailovka 2 LM (Early Turolian, MN11) Ruemkelia sp. NESIN 2013 Frunzovka 2 LM (Early Turolian, MN11) Recapitulation Ukrainian fossil insectivores 147 Table V (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Desmana cf. kormosi now LP (Early Villanyan, end of REKOVETS and PASHKOV Popovo 2 Galemys cf. kormosi MN16) 2009 (SCHREUDER, 1940) Desmana pontica now LM (Middle Turolian, Cherevychnoe 3 NESIN 2013 Ruemkelia pontica MN12) (SCHREUDER, 1940) Frunzovka 2 LM (Early Turolian, MN11) V. A. TOPACHEVSKY, NESIN and I. V. TOPA- Desmana LM (Early Turolian, Mikhailovka 2 CHEVSKY 1997; NESIN (Archaeodesmana) sp. MN11) and NADACHOWSKI 2001 Vinogradovka 1 LM (Late Turolian, MN13) NESIN 2013 Desmana (Archaeodesmana) cf. LM (Early/Middle REKOVETS and PASHKOV verestchagini Verkhnya Krynitsa 2 Turolian, MN11/MN12) 2009 TOPACHEVSKI, 1961 (in 1961a) V. O. TOPACHEVSKY, Desmana aff. Odessa, 16th Station of CHEPALIGA, NESIN, LM (Late Turolian, MN13) verestchagini Bolshoy Fontan REKOVETS and I.V. TOPACHEVSKY 1988 V. A. TOPACHEVSKY, Zhevakova Gora LP (Early Villanyan, Desmana nehringi SKORIK,CHEPALYGA (ls. 11, 15) middle of MN16) 1979 Odessa, 16th Station of LM (Late Turolian, MN13) Bolshoy Fontan Desmana cf. nehringi NESIN 2013 Zhevakova Gora (l. 4, LP (Early Villanyan, previously 15) middle of MN16) Myogale thermalis V. O. TOPACHEVSKY now Kairy EPl (Early Biharian, Q1) 1956 Desmana thermalis Kairy EPl (Early Biharian, Q1) V. O. TOPACHEVSKY Nogaisk EPl (Early Biharian, Q1) 1957b V. A. TOPACHEVSKY, Desmana thermalis Zhevakova Gora (ls. 5) EPl (Early Biharian, Q1) SKORIK, CHEPALYGA 1979 Tiligul (=Morskoe) EPl (Early Biharian, Q1) REKOVETS and Zhevakhova Gora (l. 5) EPl (Early Biharian, Q1) NADACHOWSKI 1995 B. RZEBIK-KOWALSKA, L. I. REKOVETS 148 Table V (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Desmana (Pliodesmana) LP (Early Villanyan, REKOVETS and PASHKOV moldavica PASHKOV and Verkhnya Krynitsa 1 beginning of MN16) 2009 TOPACHEVSKY, 1990 P/Pl boundary (previously Desmana (Pliodesmana) REKOVETS and PASHKOV Popovo 1 Early/Late Villanyan, cf. jalpugensis 2009 MN16/MN17) EPl (previously Late Kotlovina 2 Villanyan, MN17) Desmana (Pliodesmana) sp. NESIN 2013 EPl (previously Late Kotlovina 3 Villanyan, MN17) V. A. TOPACHEVSKY, NESIN and I. V. TOPA- EP (Early Ruscinian, Krasnopol CHEVSKY 1997; NESIN MN14) and NADACHOWSKI 2001; NESIN 2013 Desmana EP (Late Ruscinian, Kotlovina 1 (Galemodesmana) sp. MN15) EPl (previously Late Kotlovina 2 NESIN 2013 Villanyan, MN17) EPl (previously Late Kotlovina 3 Villanyan, MN17) REKOVETS and Desmana meridionalis Kryzhanovka 1 EPl (Early Biharian, Q1) NADACHOWSKI 1995 Nogaisk EPl (Early Biharian, Q1) REKOVETS and Desmana nogaica Kairy EPl (Early Biharian, Q1) NADACHOWSKI 1995 Cherevychnoe 1 EPl (Early Biharian, Q1) Karai Dubina EPl (Early Biharian, Q1) REKOVETS and Desmana gureevi Luzanovka EPl (Early Biharian, Q1) NADACHOWSKI 1995 Myogale cf. moschata now Sinyakovo EPl PIDOPLICHKO 1955 Desmana cf. moschata (LINNAEUS, 1758) V. A. TOPACHEVSKY, Morozovka 2 MPl (Steinheimian, Q3) Desmana moschata KORNIETS, SVISTUN 1975 Borisova Gora LPl MARKOVA 1987 Desmana moschata Sinyakovo EPl PIDOPLICHKO 1956 ternopolitana Recapitulation Ukrainian fossil insectivores 149 Table V (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species LP (?Villanyan) and ?EPl near Kryzhanovka (?Q1) SHEVCHENKO 1965 near Leventsovka ?LP (?Late Villanyan) Verkhnyaya Emancha MPl (Q3) MARKOVA 1980 Bolshevik 2 (l. III) EPl (Late Biharian, Q2) REKOVETS, CHEPALYGA, NESIN and SVETLITSKAYA Bolshevik 2 (l. I) MPl (Steinheimian, Q3) 1990; REKOVETS 1990 Tikhonovka 2 EPl (Q1) Bolshevik 2 (l. III) EPl (Late Biharian, Q2) Protopopovka (ls. 1 Desmana sp. EPl (Late Biharian, Q2) and 2) REKOVETS and Morozovka 2 MPl (Steinheimian, Q3) NADACHOWSKI 1995 Bolshaya MPl (Steinheimian, Q3) Kamyshevakha 1 Gunki MPl (Late Steinheimian, Q3) Bolshevik 2 (l. I) MPl (Steinheimian, Q3) V. A. TOPACHEVSKY, NESIN and I. V. TOPA- Mikhailovka 1 LM (Late Vallesian, MN10) CHEVSKY 1997; NESIN and NADACHOWSKI 2001 Desmana sp. ? near Kryzhanovka EPl ?(Q1) SHEVCHENKO 1965 Desmaninae gen. et sp. Mikhailovka 1 LM (Late Vallesian, MN10) NESIN 2013 indet. Frunzovka 2 LM (Early Turolian, MN11) Uropsilinae Desmanella cf. dubia Mikhailovka 1 LM (Late Vallesian, MN10) NESIN 2013 Desmanella sp. Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 Scalopinae Scalopini V. A. TOPACHEVSKY,NESIN and I. V. TOPACHEVSKY 1997; RZEBIK-KOWALSKA Domninoides sp. Gritsev LM (Early Vallesian, MN9) and V. O. TOPACHEVSKY 1997; NESIN and NADA- CHOWSKI 2001; NESIN 2013 RZEBIK-KOWALSKA and Proscapanus sp. Gritsev LM (Early Vallesian, MN9) V. O. TOPACHEVSKY 1997; NESIN 2013 B. RZEBIK-KOWALSKA, L. I. REKOVETS 150 Table V (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Proscapanus NESIN and Gritsev LM (Early Vallesian, MN9) (=Alloscapanus) NADACHOWSKI 2001 V. A. TOPACHEVSKY, NESIN and I. V. TOPA- Gritsev LM (Early Vallesian, MN9) ?Urotrichinae CHEVSKY 1997; NESIN and NADACHOWSKI 2001 Novoelizavetovka 3 LM (Middle Turolian, MN12) NESIN 2013 Talpinae Urotrichini Urotrichus giganteus Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 ?Scaptonyx sp. Mikhailovka 1 LM (Late Vallesian, MN10) NESIN 2013 RZEBIK-KOWALSKA and V. O. TOPACHEVSKY 1997; Urotrichini sp. 1 and sp. 2 Gritsev LM (Early Vallesian, MN9) NESIN and NADACHOWSKI 2001; NESIN 2013 Talpini Talpa gilothi Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 Talpa minor Gorishnya Vygnanka LP TATARINOV 1965b FREUDENBERG, 1914 Talpa cf. praeglacialis REKOVETS and Cherevychnoe 1 EPl (Early Biharian, Q1) KORMOS, 1930 NADACHOWSKI 1995 Talpa europaea Sinyakovo EPl PIDOPLICHKO 1955, 1956 Talpa ex gr. europaea Mikhailovka 5 LPl (Eem) AGADZHANYAN 2009 Krasnopol EP (Early Ruscinian, MN14) Kotlovina 1 EP (Late Ruscinian, MN15) EPl (previously Late Kotlovina 2 NESIN 2013 Villanyan, MN17) Talpa sp. EPl (previously Late Kotlovina 3 Villanyan, MN17) Leventsovka ?LP (?Late Villanyan) SHEVCHENKO 1965 REKOVETS and Gunki MPl (Steinheimian, Q3) NADACHOWSKI 1995 Talpa sp. ? near Kryzhanovka EPl (Q1) SHEVCHENKO 1965 Talpidae incertae sedis Desmanodon major Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 Talpidae gen. et sp. indet. Cherevychnoe 3 LM (Middle Turolian, MN12) NESIN 2013 Recapitulation Ukrainian fossil insectivores 151
Table VI Fossil Talpidae of Ukraine (mentioned without descriptions, measurements or exact localities)
Family Subfamilies Localities Age Publications Genera Species Talpidae Desmaninae alluvia (lower course of Dnieper) LPl and H V. O. TOPACHEVSKY 1957a Desmana moschata alluvia (middle course of Dnieper, Cherkasy-Mishurin Rig and MPl and LPl V. O. TOPACHEVSKY 1961b Kiev-Cherkasy) Desmana moschata Podolia and Prikarpatie (north to HTATARINOV 1970 ternopolitana Carpathian Mts) Talpinae Podolia and Prikarpatie (north to Talpa minor PTATARINOV 1970 Carpathian Mts) Podolia and Prikarpatie (north to Talpa cf. europaea Pl and H TATARINOV 1970 Carpathian Mts) alluvia (lower course of Dnieper) LPl and H V. O. TOPACHEVSKY 1957a alluvia (middle course of Dnieper, Cherkasy-Mishurin Rig and MPl and LPl V. O. TOPACHEVSKY 1961b Talpa europaea Kiev-Cherkasy) Podolia and Prikarpatie (north to Pl and H TATARINOV 1970 Carpathian Mts)
The fossil mole Proscapanus GAILLARD, 1899 (Scalopini) mentioned by TOPACHEVSKY (in RZEBIK-KOWALSKA and TOPACHEVSKY 1997) from Gritsev is known from several European localities (MN4-MN11), among others from Slovakia and the Re- public of Moldova, therefore it is also probable in Ukraine. The combination Proscapanus (Alloscapanus) cited by NESIN and NADACHOWSKI (2001) is incorrect and Alloscapanus BAUDELOT, 1968 is not a valid genus because it is a junior synonym of Proscapanus GAIL- LARD, 1899 (VAN DEN HOEK OSTENDE and FURIÓ 2005). The fossil Talpa minor, largely distributed in Europe in the Pliocene and Pleistocene, was noted by TATARINOV (1965b) from the Late Pliocene and the recent T. europaea LIN- NAEUS, 1758 by TOPACHEVSKY (1961b) from the Middle and Late Pleistocene. Their fos- sils were never thoroughly described but both species are very probably present in this part of Europe. On the other hand T. cf. praeglacialis mentioned by REKOVETS and NADA- CHOWSKI (1995) from Cherevychnoe (Late Biharian?) most probably represents a recent T. europaea. According to many authors (KRETZOI 1938, HELLER 1958, VON KOENIGSWALD 1970, RABEDER 1972 and others) T. praeglacialis, T. fossilis and the recent T. europaea are similar in size and morphology and they should be considered as one species. The names T. fossilis (= T. praeglacialis) indicate only a chronospecies. B. RZEBIK-KOWALSKA, L. I. REKOVETS 152
II.3. Family Dimylidae SCHLOSSER, 1887 (Table VII) Comments. So far the Dimylidae family known from Europe and the Middle East (MP27-MN11) is represented in Ukraine only in one locality and according to TOPACHEVSKY (in RZEBIK-KOWALSKA and TOPACHEVSKY 1997) by one genus, Plesio- dimylus GAILLARD, 1897. This form is characterised by less specialized dentition, most probably adapted to an insectivorous diet. It is the most widespread (found in more than 70 localities) of all European Dimylidae taxa and is known from the Middle and Late Mio- cene (MN4 to MN11), among others from Poland (RZEBIK-KOWALSKA 1996), Slovakia (FEJFAR and SABOL 2005) and Hungary (HIR and MÉSZÁROS 2002, PRIETO et al. 2012). According to SCHMIDT-KITTLER (1973), the family is related to Soricidae (absence of a zygomatic arch) rather than to the Erinaceidae (RZEBIK-KOWALSKA 2009). However, a closer relationship between Dimylidae and Talpidae is corroborated by some fossil finds from Anatolia (VAN DEN HOEK OSTENDE 1995).
Table VII
Dimylidae of Ukraine (mentioned without descriptions or measurements)
Family Genus Locality Age Publications Species
Dimylidae
V. A. TOPACHEVSKY,NESIN and I. V. TOPACHEVSKY 1997; RZEBIK-KOWALSKA Plesiodimylus sp. Gritsev LM (Early Vallesian, MN9) and V. O. TOPACHEVSKY 1997; NESIN and NADA- CHOWSKI 2001; NESIN 2013
II.4. Family Heterosoricidae VIRET and ZAPFE, 1951 (Tables VIII-IX) Comments. So far only one taxon was found in Ukraine. Dinosorex cf. zapfei ENGES- SER, 1975 (TOPACHEVSKY et al. 1997) was later described as a new species, D. grycivensis RZEBIK-KOWALSKA and TOPACHEVSKY, 1997. The genus (with numerous species) is known from the Late Oligocene (MP29) to the Late Miocene (MN10) from many Euro- pean countries. D. grycivensis was also found in Poland, the Republic of Moldova and is largely represented in Spain (FURIÓ et al. 2015), thus opening serious questions about the real extent of its distribution in the past. Recapitulation Ukrainian fossil insectivores 153 Table VIII Heterosoricidae of Ukraine (published with descriptions, measuremets, localities and geo- logical ages)
Family Genus Locality Age Publication Species
Heterosoricidae
Dinosorex grycivensis RZEBIK-KOWALSKA and RZEBIK-KOWALSKA and Gritsev LM (Early Vallesian, MN9) V. O. TOPACHEVSKY 1997 TOPACHEVSKY, 1997
Table IX Heterosoricidae of Ukraine (mentioned without descriptions or measurements)
Family Genus Localities Age Publications Species
Heterosoricidae
?Dinosorex zapfei Gritsev LM (Early Vallesian, MN9) NESIN and NADACHOWSKI 2001 ENGESSER, 1975
V. A. TOPACHEVSKY, NESIN Dinosorex cf. zapfei Gritsev LM (Early Vallesian, MN9) and I. V. TOPACHEVSKY 1997
Dinosorex grycivensis Gritsev LM (Early Vallesian, MN9) NESIN 2013
II.5. Family Soricidae FISCHER, 1814 (Tables X-XII) Comments. In Ukraine Miosorex sp. was cited for the first time by TOPACHEVSKY et al. (1997) from Cherevychnoe dated to MN12. If the identification is correct, these remains are the youngest in Europe because this common Miocene genus, known from almost all European countries (from Portugal to the Republic of Moldova), was dated so far to MN3-MN11 (RZEBIK-KOWALSKA 2009). Its presence in Ukraine (although in older lo- calities, MN11) was confirmed by RZEBIK-KOWALSKA and NESIN (2010) and RZE- BIK-KOWALSKA and REKOVETS (in press). Pachyura sp. mentioned in TOPACHEVSKY (1956) probably represents the genus Pae- nelimnoecus (see REUMER 1984). In Europe Paenelimnoecus is known from the early B. RZEBIK-KOWALSKA, L. I. REKOVETS 154 Middle Miocene (MN3) to the Late Pliocene (MN16). In 2006 it was transferred by FEJFAR,STORCH, and TOBIEN from the subfamily Allosoricinae to the newly created by them subfamily Paenelimnoecinae. In 1988 TOPACHEVSKY et al., cited it from Cher- evychnoe (MN12). Its presence in Ukraine is probable because it is also known from neighboring countries such as Poland, Hungary, Slovakia and Romania. The only species correctly described from Ukraine, “Paenelimnoecus” repenningi, belongs probably to an- other genus (see RZEBIK-KOWALSKA 2009). Anourosoricini are represented by three or four genera (Amblycoptus, Crusafontina, Paranourosorex and potentially Anourosoricodon) in Ukraine (RZEBIK-KOWALSKA 1998). Paranourosorex sp. was listed for Ukraine from the late Early Pliocene (MN15) by TOPACHEVSKY et al. (1997) and NESIN and NADACHOWSKI (2001), however without any description or measurements. Because it was described from nearby Poland (early Early Pliocene, MN14, RZEBIK-KOWALSKA 1975) and cited from Slovakia (MN15, FEJFAR and SABOL 2005) it is also expected in neighboring territories. It was also found in Asia (Ka- zakhstan, China, between MN12 and MN14, STORCH and ZAZHIGIN 1996). On the other hand, Anourosorex sp. cited by TOPACHEVSKY (1962b) from the “Middle Pliocene” of Kamyanske was later identified as Anourosoricodon pidoplitschkoi TOPACHEVSKY, 1966. So far the latter was never found in any other country. In the papers of TOPACHEVSKY et al. (1997) and RZEBIK-KOWALSKA and TOPACHEVSKY (1997), TOPACHEVSKY supposed that Anourosoricodon may be a synonym of Crusafontina. However, this opinion as well as the suggested presence in Gritsev of such taxa as Lan- thanotherium, Amphechinus, Domninoides, Proscapanus and Plesiodimylus were never confirmed by morphological study. The paper on Gritsev (RZEBIK-KOWALSKA and TOPACHEVSKY 1997) had no continuation because of the death of professor V. O. TOPACHEVSKY. Blarina ukrainica described by PIDOPLICHKO in 1955 was later included into Beremendia (GUREEV 1971). CROCHET and MICHAUX (1981) mentioned Beremendia cf. ucrainica from the Middle Pleistocene of France and RZEBIK-KOWALSKA (1998) placed it in Bere- mendia fissidens (Beremendiini). In general, Sulimskia is also expected in Ukraine because it was described from Poland (MN15) and later found in other European countries, e.g. in Slovakia and Hungary (in lo- calities dated between MN14 and the Pliocene/Pleistocene boundary, RZEBIK-KOWALSKA 1998). It is also known from western Siberia, the Transbaikalia region (Late Pliocene, STORCH et al. 1998), inner Mongolia (QIU and STORCH 2000) and China (MN14-MN15, QIU and STORCH 2005). However, its occurrence in the Late Miocene (MN11) localities (cited by TOPACHEVSKY et al. 1997, NESIN and NADACHOWSKI 2001 and NESIN 2013 without any description) is less probable. The presence of Neomysorex mentioned in Ukraine by TOPACHEVSKY et al. (1997) and NESIN and NADACHOWSKI (2001) (MN15) was later confirmed by RZEBIK-KOWALSKA and NESIN (2010) and RZEBIK-KOWALSKA and REKOVETS (in press). Ukrainian remains found in Late Miocene (MN11-MN12) localities are the oldest known so far. It is also known from the Early Pliocene (MN14-MN14/MN15) of Poland (RZEBIK-KOWALSKA 1981). Recapitulation Ukrainian fossil insectivores 155
The Early Pleistocene Neomys newtoni (Neomyini) was found in Ukraine only once at Medzhybozh (Q3) (RZEBIK-KOWALSKA and REKOVETS, in press). It is known from neighboring Poland and Romania. It was also cited by ZAITSEV and BARYSHNIKOV (2002) from the northern Caucasus and from the Early and Middle Pleistocene of Europe (RZEBIK-KOWALSKA 1998).
The recent Neomys fodiens (PENNANT, 1771) was cited by TOPACHEVSKY (1961b) in Middle and Late Pleistocene alluvial deposits. So far it has been found in almost all Euro- pean countries (RZEBIK-KOWALSKA 1998). Today its large range over the entire northern and central Palaearctic includes Ukrainian territory.
Episoriculus cited from Gritsev by NESIN and NADACHOWSKI (2001) is improbable in Ukraine. Species of this genus now live in southeastern Asia. If remains from Gritsev really belong to the Neomyini, they could be representatives of Asoriculus. It was men- tioned by TOPACHEVSKY in RZEBIK-KOWALSKA and TOPACHEVSKY (1997) and later confirmed by RZEBIK-KOWALSKA and NESIN (2010) and RZEBIK-KOWALSKA and REK- OVETS (in press). Its remains from Frunzovka 2 (MN11) are the oldest known so far. This genus was widely distributed in Europe from MN13 to the Early Pleistocene (RZE- BIK-KOWALSKA 2009) and was also collected in Asia Minor (MN14-MN15, STORCH et al. 1998). Both species of Crocidura (C. leucodon and C. suaveolens) are possible as fossils in Ukraine because they live there today.
On the other hand, the presence of Suncus is less probable although TOPACHEVSKY (1957b, 1965) cited it (without description) from the Late Pliocene of Kairy. So far there are no unambiguous published records of fossil European Suncus. The single specimen (i1) found in Petralona (Greece, SICKENBERG 1971) could represent this genus but its as- signment is uncertain.
Sorex arcticus KERR, 1792 described by MEZHZHERIN (1972) from the Late Pleisto- cene of alluvial deposits of Desna and Dnieper rivers is quite impossible in the area of Ukraine. Today, the species lives in Canada (from Yukon to Quebec) and in the northern part of the USA.
Sorex praearaneus praearaneus described by MEZHZHERIN in the same paper (1972) is now placed in the subgenus S.(Drepanosorex) praearaneus. It appeared in the Early Pleistocene (former MN17) and survived probably until the Late Pleistocene. It also lived in Poland, Hungary (type locality), Slovakia and Romania (RZEBIK-KOWALSKA 2009).
Remains of the recent Sorex araneus LINNAEUS, 1758 described by PIDOPLICHKO (1955, 1956) and TOPACHEVSKY (1961b) from the Early and Middle Pleistocene alluvial deposits of Ukraine could have also belonged (especially older Middle Pleistocene re- mains) to fossil S. subaraneus HELLER, 1958. Now it is impossible to say when S. sub- araneus disappeared and S. araneus appeared on the European continent (RZEBIK-KOWAL- SKA 2009). Only a revision of all European fossils of S. subaraneus and S. araneus could help in understanding this problem as well as the question of ancestry of the recent species. B. RZEBIK-KOWALSKA, L. I. REKOVETS 156 Table X Fossil Soricidae of Ukraine (published with descriptions, measuremets, localities and geo- logical ages)
Family Subfamilies Tribes Localities Age Publications Genera Species Soricidae Crocidosoricinae Miosorex grivensis LM (Early Turolian, RZEBIK-KOWALSKA and Popovo 3 (DEPÉRET, 1892) MN11) REKOVETS in press LM (Early Turolian, RZEBIK-KOWALSKA and Mikhailovka 2 MN11) NESIN 2010 cf. Miosorex sp. LM (Early/Middle RZEBIK-KOWALSKA and Verkhnya Krynitsa 2 Turolian, MN11/MN12) REKOVETS in press Paenelimnoecinae „Paenelimnoecus” repenningi LM (Early Turolian, RZEBIK-KOWALSKA and Frunzovka 2 (BACHMAYER and MN11) NESIN 2010 WILSON, 1970) Soricinae Anourosoricini Anourosoricodon EP (Late Ruscinian, V. O. TOPACHEVSKY 1966 pidoplitschkoi Kamenskoe MN15) UREEV TOPACHEVSKY, 1966 G 1971, 1979 LM (Early Turolian, Mikhailovka 2 Amblycoptus oligodon MN11) RZEBIK-KOWALSKA and ORMOS ESIN K , 1926 LM (Early Turolian, N 2010 Frunzovka 2 MN11) Amblycoptus sp. or Odessa, 16th Station of RZEBIK-KOWALSKA and LM (Late Turolian, MN13) Kordosia sp. Bolshoy Fontan NESIN 2010 LM (Late Vallesian, Mikhailovka 1 Crusafontina kormosi MN10) ZEBIK OWALSKA ACHMAYER R -K and (B and ESIN ILSON LM (Early Turolian, N 2010 W , 1970) Frunzovka 2 MN11) LM (Early/Middle RZEBIK-KOWALSKA and Crusafontina cf. kormosi Verkhnya Krynitsa 2 Turolian, MN11/MN12) REKOVETS in press Recapitulation Ukrainian fossil insectivores 157 Table X (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Beremendiini LP (Early Villanyan, end of Popovo 2 MN16) P/Pl boundary (previously Beremendia fissidens RZEBIK-KOWALSKA and Popovo 1 Early/Late Villanyan, (PÉTENYI, 1864) REKOVETS in press MN16/MN17) EPl (previously Late Popovo 0 Villanyan, MN17) Chortkov EPl (Early Biharian, Q1) PIDOPLICHKO 1955 Blarina ucrainica PIDOPLICHKO, 1955 Gorishnya Vygnanka LP now ATARINOV Verkhnyaya T 1958 Beremendia fissidens ?EPl Vygnanka cf. Beremendia minor LP (Early Villanyan, RZEBIK-KOWALSKA and RZEBIK-KOWALSKA, Verkhnya Krynitsa 1 beginning of MN16) REKOVETS in press 1976 Blarinellini LM (Early/Middle Petenyia dubia Verkhnya Krynitsa 2 ZEBIK OWALSKA ACHMAYER Turolian, MN11/MN12 R -K and B and EKOVETS ILSON R in press W , 1970 Lobkove LM (Middle Turolian, MN12) RZEBIK-KOWALSKA and Petenyia cf. dubia Frunzovka 2 LM (Early Turolian, MN11) NESIN 2010 Petenyia hungarica LP (Early Villanyan, RZEBIK-KOWALSKA and Verkhnya Krynitsa 1 KORMOS, 1934 beginning of MN16) REKOVETS in press Odessa, 16th Station of RZEBIK-KOWALSKA and ?Petenyia sp. LM (Late Turolian, MN13) Bolshoy Fontan NESIN 2010 Blarinini Mafia cf. dehneli RZEBIK-KOWALSKA and Mikhailovka 2 LM (Early Turolian, MN11) (KOWALSKI, 1956) NESIN 2010 Neomyini Neomysorex alpinoides LM (Early/Middle RZEBIK-KOWALSKA and Verkhnya Krynitsa 2 (KOWALSKI, 1956) Turolian, MN11/MN12) REKOVETS in press B. RZEBIK-KOWALSKA, L. I. REKOVETS 158 Table X (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species LM (Early Turolian, Mikhailovka 2 ?Neomysorex cf. MN11) RZEBIK-KOWALSKA and alpinoides NESIN 2010 Frunzovka 2 LM (Early Turolian, MN11)
Asoriculus cf. th Odessa, 16 Station of RZEBIK-KOWALSKA and gibberodon (PETÉNYI, LM (Late Turolian, MN13) Bolshoy Fontan NESIN 2010 1864) RZEBIK-KOWALSKA and Asoriculus sp. Frunzovka 2 LM (Early Turolian, MN11) NESIN 2010 Popovo 3 LM (Early Turolian, MN11) RZEBIK-KOWALSKA and cf. Asoriculus sp. LM (Early/Middle EKOVETS Verkhnya Krynitsa 2 R in press Turolian, MN11/MN12) Neomys newtoni RZEBIK-KOWALSKA and Medzhybozh MPl (Q3) HINTON, 1911 REKOVETS in press Soricini Sorex pseudoalpinus RZEBIK-KOWALSKA and Mikhailovka 2 LM (Early Turolian, MN11) RZEBIK-KOWALSKA, 1991 NESIN 2010 Sorex praearaneus Nogaisk EPl (Early Biharian, Q1) praearaneus now MEZHZHERIN 1972 S. (Drepanosorex) Ozernoe MPl praearaneus KORMOS, 1934 MEZHZHERIN and SVISTUN Novogrod Severskii LPl (Late Weichselian) Sorex araneus 1968 LINNAEUS, 1758 Mikhailovka 5 LPl (Eem) AGADZHANYAN 2009 Sorex arcticus KERR, 1792 Novogrod Severskii LPl (Late Weichselian) MEZHZHERIN 1972
EZHZHERIN Sorex minutus Khrenniki LPl M 1972 INNAEUS L , 1766 Mikhailovka 5 LPl (Eem) AGADZHANYAN 2009 Zelceina cf. RZEBIK-KOWALSKA and podlesicensis Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2010 RZEBIK-KOWALSKA, 1990 LM (Early/Middle RZEBIK-KOWALSKA and Zelceina sp. Verkhnya Krynitsa 2 Turolian, MN11/MN12) REKOVETS in press Recapitulation Ukrainian fossil insectivores 159 Table XI Fossil Soricidae of Ukraine (mentioned without descriptions or measurements)
Family Subfamilies Tribes Localities Age Publications Genera Species Soricidae Crocidosoricinae ?Miosorex sp. Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 V. A. TOPACHEVSKY, NESIN and I. V. TOPA- LM (Middle Turolian, Miosorex sp. Cherevychnoe 3 CHEVSKY 1997; NESIN MN12) and NADACHOWSKI 2001; NESIN 2013 ?Paenelimnoecinae or ?Crocidurinae Pachyura sp. V. O. TOPACHEVSKY Kairy EPl (Early Biharian, Q1) (?Paenelimnoecus or 1956 ?Suncus) Paenolimnoecinae „Paenelimnoecus” Frunzovka 2 LM (Early Turolian, MN11) NESIN 2013 repenningi LM (Middle Turolian, Novoelizavetovka 3 NESIN 2013 MN12) V. A. TOPACHEVSKY, Paenelimnoecus sp. NESIN and I. V. LM (Middle Turolian, Cherevychnoe 3 TOPACHEVSKY 1997; MN12) NESIN and NADACHOWSKI 2001 LM (Early Turolian, Novoelizavetovka 2 MN11) “Paenelimnoecus” sp. NESIN 2013 LM (Middle Turolian, Cherevychnoe 3 MN12) Soricinae Anourosoricini Amblicoptini (should be V. A. TOPACHEVSKY, Amblycoptini, NESIN and I. V. TOPA- Gritsev LM (Early Vallesian, MN9) now CHEVSKY 1997; NESIN Anourosoricini) and NADACHOWSKI 2001 B. RZEBIK-KOWALSKA, L. I. REKOVETS 160 Table XI (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species
Anourosorex sp. Kamenskoe EP (Late Ruscinian, MN15) V. O. TOPACHEVSKY 1962b (now Anourosoricodon)
RZEBIK-KOWALSKA and Anourosoricodon LM (Early Vallesian, Gritsev V. O. TOPACHEVSKY (=?Crusafontina) MN9) 1997; NESIN 2013
V. A. TOPACHEVSKY, NESIN and I. V. Anourosoricodon LM (Early Vallesian, Gritsev TOPACHEVSKY 1997; (=Crusafontina) MN9) NESIN and NADACHOWSKI 2001
V. A. TOPACHEVSKY, Amblicoptus sp. (should be LM (Early Turolian, Mikhailovka 2 NESIN and I. V. Amblycoptus) MN11) TOPACHEVSKY 1997 Mikhailovka 2 LM (Early Turolian, MN11) Amblycoptus oligodon NESIN 2013 Frunzovka 2 LM (Early Turolian, MN11)
LM (Early Turolian, NESIN and NADACHOWSKI Novoelizavetovka 2 MN11) 2001; NESIN 2013
Novoelizavetovka 3 LM (Middle Turolian, NESIN 2013 MN12) Amblycoptus sp. Cherevychnoe 3 V. O. TOPACHEVSKY, Odessa, 16th Station CHEPALYGA, NESIN, LM (Late Turolian, MN13) of Bolshoy Fontan REKOVETS and I.V. TOPACHEVSKY 1988
Amblycoptus sp. or Odessa, 16th Station LM (Late Turolian, MN13) NESIN 2013 Kordosia sp. of Bolshoy Fontan Mikhailovka 1 LM (Late Vallesian, MN10) Crusafontina kormosi NESIN 2013 Frunzovka 2 LM (Early Turolian, MN11)
V. A. TOPACHEVSKY, NESIN and I. V. TOPA- Paranourosorex sp. Kotlovina 1 ?EP (Late Ruscinian, MN15) CHEVSKY 1997; NESIN and NADACHOWSKI 2001 Recapitulation Ukrainian fossil insectivores 161 Table XI (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Beremendiini PIDOPLICHKO 1956; Chortkov EPl Early Biharian, Q1) Blarina ucrainica TATARINOV 1958 now Gorishnya Vygnanka LP TATARINOV 1965b Beremendia fissidens Verkhnyaya Vygnanka ?EPl TATARINOV 1958 V. A. TOPACHEVSKY, SKORIK, CHEPALYGA Zhevakova Gora (l. 9) EPl (Early Biharian, Q1) 1979; REKOVETS and NADACHOWSKI 1995 V. A. TOPACHEVSKY,NESIN andI.V.TOPACHEVSKY 1997; Kotlovina 1 ?EP (Late Ruscinian, MN15) Beremendia sp. NESIN and NADACHOWSKI 2001 EPl (previously Late Kotlovina 2 Villanyan, MN17) NESIN 2013 EPl (previously Late Kotlovina 3 Villanyan, MN17) Blarinini Mafia cf. dehneli Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 Novoelizavetovka 2 LM (Early Turolian, MN11) Sulimskia sp. Novoelizavetovka 3 LM (Middle Turolian, MN12) NESIN 2013 Cherevychnoe 3 LM (Middle Turolian, MN12) V. A. TOPACHEVSKY, ?Sulimskia sp. Mikhailovka 2 LM (Early Turolian, MN11) NESIN and I. V. TOPACHEVSKY 1997 NESIN and cf. Sulimskia sp. Novoelizavetovka 2 LM (Early Turolian, MN11) NADACHOWSKI 2001 NESIN and Blarinini gen. et sp. indet. Novoukrainka 1 LM (Late Turolian, MN13) NADACHOWSKI 2001 Blarinellini Petenyia cf. dubia Frunzovka 2 LM (Early Turolian, MN11) NESIN 2013 Odessa, 16th Station ?Petenyia sp. LM (Late Turolian, MN13) NESIN 2013 of Bolshoy Fontan B. RZEBIK-KOWALSKA, L. I. REKOVETS 162 Table XI (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Neomyini
?Neomysorex cf. Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 alpinoides Frunzovka 2 LM (Early Turolian, MN11) V. A. TOPACHEVSKY, NESIN and I. V. TOPA- Neomysorex sp. Kotlovina 1 EP (late Ruscinian, MN15) CHEVSKY 1997; NESIN and NADACHOWSKI 2001
Episoriculus sp. V. A. TOPACHEVSKY, NESIN and I. V. TOPA- now Gritsev LM (Early Vallesian, MN9) CHEVSKY 1997; NESIN Asoriculus sp. and NADACHOWSKI 2001 Odessa, 16th Station Asoriculus cf. gibberodon LM (Late Turolian, MN13) NESIN 2013 of Bolshoy Fontan Gritsev LM (Early Vallesian, MN9) Asoriculus sp. NESIN 2013 Frunzovka 2 LM (Early Turolian, MN11) RZEBIK-KOWALSKA and ?Asoriculus sp. Gritsev LM (Early Vallesian, MN9 V. O. TOPACHEVSKY 1997 Soriculini gen. et sp. indet. NESIN and now Novoukrainka 1 LM (Late Turolian, MN13) NADACHOWSKI 2001 Neomyini Soricini Sorex pseudoalpinus Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 Chortkov EPl (Early Biharian, Q1) PIDOPLICHKO 1955, 1956 Sorex araneus MEZHZHERIN and Novogrod Severskii LPl (Weichselian) SVISTUN 1968 Sorex praearaneus now ARKOVA Sorex (Drepanosorex) Gunki MPl (Steinheimian, Q3) M 1980 praearaneus Sorex macropygmaeus MILLER, 1901 now Chortkov EPl (Early Biharian, Q1) PIDOPLICHKO 1955, 1956 Sorex caecutiens LAXMANN, 1788 Recapitulation Ukrainian fossil insectivores 163 Table XI (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Sorex aff. runtonensis REKOVETS and Cherevychnoe 1 EPl (Early Biharian, Q1) HINTON, 1911 NADACHOWSKI 1995 Sorex cf. minutus Mikhailovka 5 LPl (Eem) AGADZHANYAN 2009 Bolshevik 1 MPl SHEVCHENKO 1965 Srednyaya Cave (near Pl TATARINOV 1965a Nizhnee Krivche) Verkhnyaya Emancha MPl (Q3) MARKOVA 1980 Gunki MPl (Steinheimian, Q3) Chernyanka LPl (Eem) MARKOVA 1985 Bolshevik 2 (l. III) EPl (Late Biharian, Q2) REKOVETS, CHEPALYGA, NESIN and Bolshevik 2 (l. I) MPl (Steinheimian, Q3) SVETLITSKAYA 1990; REKOVETS 1990 REKOVETS and STARKIN Anetovka 2 LPl (Late Weichselian) 1990 Chortkov EPl (Early Biharian, Q1) Tarkhankut EPl (Early Biharian, Q1) Kairy EPl (Early Biharian, Q1) Sorex sp. Karai Dubina EPl (Early Biharian, Q1) Luzanovka EPl (Early Biharian, Q1) Bolshevik 2 (ls. II, III) EPl (Late Biharian, Q2) Protopopovka ( l. 2) Epl (Late Biharian, Q2) Tikhonovka 1 EPl (Late Biharian, Q2) REKOVETS and Bolshevik 2 (l. I) MPl (Steinheimian, Q3) NADACHOWSKI 1995 Morozovka 2 MPl (Steinheimian, Q3) Bolshaya MPl (Steinheimian, Q3) Kamyshevakha 1 Demidovka MPl (Steinheimian, Q3) Gunki MPl (Steinheimian, Q3) Medzhybozh MPl (Steinheimian, Q3) Matveevka MPl (Steinheimian, Q3) EP (Late Ruscinian, REKOVETS and PASHKOV Kamenskoe MN15) 2009 Mikhailovka 4 MPl (Steinheimian, Q3) AGADZHANYAN 2009 Mikhailovka 5 LPl (Eem) Zelceina cf. podlesicensis Mikhailovka 2 LM (Early Turolian, MN11) NESIN 2013 B. RZEBIK-KOWALSKA, L. I. REKOVETS 164 Table XI (continuation)
Family Subfamilies Tribes Localities Age Publications Genera Species Crocidurinae Crocidura sp. [near Srednyaya Cave (near C. leucodon (HERMANN, Pl TATARINOV 1965a Nizhnee Krivche) 1780)] REKOVETS and Crocidura sp. Tarkhankut EPl (Early Biharian, Q1) NADACHOWSKI 1995 V. O. TOPACHEVSKY Suncus sp. Kairy EPl (Early Biharian, Q1) 1957b near Leventsovka LP (Villanyan) SHEVCHENKO 1965 V. A. TOPACHEVSKY,KOR- Morozovka 2 MPl (Steinheimian, Q3) NIETS and SVISTUN 1975 Borisova Gora LPl MARKOVA 1987 V. A. TOPACHEVSKY, Zhevakova Gora (ls. LP (Early Villanyan, SKORIK and CHEPALYGA 11, 15) middle of MN16) 1979
Zhevakova Gora (l. 4, LP (Early Villanyan, ESIN previously 15) middle of MN16) N 2013 V. O. TOPACHEVSKY, Odessa, 16th Station CHEPALYGA, NESIN, MN13 of Bolshoy Fontan REKOVETS and I.V. TOPACHEVSKY 1988 V. A. TOPACHEVSKY, Soricidae gen. et sp. indet. Orekhovka LM (MN13) NESIN and PRISYAZHNYUK 1990 REKOVETS and Tiligul (=Morskoe) EPl (Early Biharian, Q1) NADACHOWSKI 1995 LM (Early Turolian, Popovo 3 MN11) LM (Early/Middle Verkhnya Krynitsa 2 Turolian, MN11/MN12) LP (Early Villanyan, REKOVETS and PASHKOV Verkhnya Krynitsa 1 beginning of MN16) 2009 LP (Early Villanyan, end Popovo 2 of MN16 P/Pl boundary (previously Popovo 1 Early/Late Villanyan, MN16/MN17 Recapitulation Ukrainian fossil insectivores 165 Table XII Fossil Soricidae of Ukraine (mentioned without descriptions, measurements or exact lo- calities) Subfamilies Tribes Localities Age Publications Genera Species Soricidae Soricinae Beremendiini middle course of EP (Early Ruscinian, Blarina (?) sp. ? SHEVCHENKO 1965 Kychurgan river MN14) Blarina ucrainica Podolia and Prikarpatie now P, EPl TATARINOV 1970 (north to Carpathian Mts) Beremendia fissidens Soricini Sorex praearaneus alluvia of Desna and Pl MEZHZHERIN 1972 praetetragonurus Dnieper rivers Sorex arcticus alluvia of Desna and LPl MEZHZHERIN 1972 (with description) Dnieper rivers alluvia (middle course of Dnieper, V. O. TOPACHEVSKY MPl and LPl Cherkasy-Mishurin Rig, 1961b Kiev-Cherkasy) Sorex araneus Podolia and Prikarpatie EPl and H TATARINOV 1970 (north to Carpathian Mts) MEZHZHERIN and alluvia of Dnieper river LPl, EH SVISTUN 1968 Sorex araneus praetetragonurus MEZHZHERIN and alluvia of Dnieper river LPl, EH MEZHZHERIN and SVISTUN 1966, 1968 SVISTUN, 1966 Podolia and Prikarpatie Sorex cf. araneus Pl TATARINOV 1970 (north to Carpathian Mts) Podolia and Prikarpatie Sorex minutus HTATARINOV 1970 (north to Carpathian Mts) Sorex macropygmaeus Podolia and Prikarpatie now EPl TATARINOV 1970 (north to Carpathian Mts) Sorex caecutiens Srednyaya Cave (near Pl TATARINOV, 1965a Nizhnee Krivche) Sorex sp. Podolia and Prikarpatie Pl and H TATARINOV, 1970 (north to Carpathian Mts) B. RZEBIK-KOWALSKA, L. I. REKOVETS 166 Table XII (continuation)
Subfamilies Tribes Localities Age Publications Genera Species basins of Salcha and Kagul EP (Late Ruscinian, Sorex sp. ? SHEVCHENKO 1965 rivers MN15) Neomyini alluvia (middle course of MPl and LPl V. O. TOPACHEVSKY, 1961b Neomys fodiens Dnieper, Kiev-Cherkasy) (PENNANT, 1771) Podolia and Prikarpatie HTATARINOV, 1970 (north to Carpathian Mts) Podolia and Prikarpatie Neomys sp. HTATARINOV, 1970 (north to Carpathian Mts) Crocidurinae alluvia (middle course of MPl and LPl V. O. TOPACHEVSKY, 1961b Dnieper, Kiev-Cherkasy) Crocidura leucodon Podolia and Prikarpatie HTATARINOV, 1970 (north to Carpathian Mts) alluvia (middle course of V. O. TOPACHEVSKY, MPl and LPl Crocidura suaveolens Dnieper, Kiev-Cherkasy) 1961b (PALLAS, 1811) Podolia and Prikarpatie HTATARINOV, 1970 (north to Carpathian Mts) Podolia and Prikarpatie Crocidura sp. Pl and H TATARINOV, 1970 (north to Carpathian Mts)
III. CONCLUSIONS
The list of the Ukrainian fossil insectivore mammals (Eulipotyphla) is still incomplete and some taxa cited in this list are wrongly identified or improbable in this country. The re- capitulation and arrangement of data present in the paper together with comments concern- ing uncertain forms should help in studies of this group of mammals in Ukraine. Each new well described form enlarges the knowledge on these mammals in less explored Eastern Europe.
REFERENCES
AGADZHANYAN A.K. 2009. Melkie mlekopitayushchie pliotsen- pleistotsena Russko0 Ravniny. Trudy Pale- ontologicheskogo Instituta, 289: 3-676. (In Russian). AYMARD A. 1850. Restes de Mammifères fossils recueillis dans le calcaire miocène des environs du Puy. An- nales de la Société d Agriculture, Sciences, Arts et Commerce du Puy, 14: 104-114. Recapitulation Ukrainian fossil insectivores 167
BACHMAYER F., WILSON R. 1970. Die Fauna der altpliozänen Höhlen- und Spalten-Füllungen bei Koh- fidisch, Burgenland (Österreich). Annalen des Naturhistorischen Museums, 74: 533-587. BARRET-HAMILTON G.E.H. 1900. Note on the common hedgehog (Erinaceus europaeus LINNAEUS) and its subspecies or local variations. The Annals and Magazine of Natural History, 5: 360-368. BAUDELOT S. 1968. A propos des Talpidés (Insectivores) de Sansan. Compte Rendu Sommaire des Séances de la Société Géologique de France, 4: 124-126. CROCHET J.-Y., MICHAUX J. 1981. Une faune de vertébrés du Pléistocène moyen sur le Causse du Larzac: Saint-Sauver, près Nant (Aveyron). Paléobiologie Continentale, 12: 131-143. DEPÉRET C. 1892. La faune de mammifères miocènes de La Grive Saint-Alban (Isère) et de quelques autres localités du Bassin du Rhône. Documents nouveaus et revision générale. Archives du Muséum dHistoire Naturelle de Lyon, 5: 1-95. ENGESSER B. 1975. Revision der europäischen Heterosoricinae (Insectivora, Mammalia). Eclogae Geologi- cae Helvetiae, 68: 649-671. ENGESSER B. 1980. Insectivora und Chiroptera (Mammalia) aus dem Neogen der Türkei. Schweizerische Paläontologische Abhandlungen, 102: 47-149. FEJFAR O., SABOL M. 2005. Czech Republic and Slovak Republic. [In:] L. W. VAN DEN HOEK OSTENDE, C. S. DOUKAS, J. W. F. REUMER (eds) – The fossil record of the Eurasian Neogene insectivores (Erinaceomorpha, Soricomorpha, Mammalia), Part. I. Scripta Geologica Special Issue, 5: 51-60. FILHOL H. 1888. Sur un noveau genre d’insectivore. Bulletin dela Societé philomatique de Paris, serie 7, 12: 24-25. FISCHER G. 1813-1814. Zoognosia tabulis synopticis illustrata. NICOLAI SERGEIDIS VSEVOLOZSKY, Mos- cow. 3 volumes: 3-1814 pp. FREUDENBERG W. 1914. Die Säugetiere des älteren Quartärs von Mitteleuropa mit besonderer Berücksichti- gung der Fauna von Hundsheim und Deutschaltenburg in Niederösterreich etc. Geologische und palaeon- tologische Abhandlungen (Neue Folge), 12: 1-219. FURIÓ M., PRIETO J., VAN DEN HOEK OSTENDE L. 2015. Three millions years of Terror Shrew (Dinosorex, Eulipotyphla, Mammalia) in the Miocene of the Vallès-Penedès Basin (Barcelona, Spain). Comptes Rendus Palevol, 14: 111-124. GAILLARD C. 1897. Nouveau genre dinsectivore du Miocène de La Grive-Saint-Alban (Isère). Comptes- rendus hebdomadaires des séances de lAcademie des sciences Paris, 124: 1248-1250. GAILLARD C. 1899. Mammifères miocènes nouveaux ou peu connus de La Grive-St.-Alban. Archives du Muséum dHistoire Naturelle de Lyon, 7: 1-79. GIBERT J. 1975. New Insectivores from the Miocene of Spain I. and II. Proceedings. Koninklijke Nederlandse Akademie van Wetenschappen, Section B, 78: 108-133. GMELIN S. G. 1770. De Capra saiga et Erinaceo aurito. Novi Commentarii Academiae Scientiarum Imperialis Petropolitanae, 14: 512-524. GREEN M. 1956. The lower Pliocene Ogallala-Wolf Creek vertebrate fauna, South Dakota. Journal of Paleon- tology, 30: 146-159. GUREEV A.A. 1971. Zemlero0ki (Soricidae) fauny mira. Publishing House Nauka, Leningrad. (In Russian). GUREEV A.A. 1979. Fauna SSSR, mlekopitayushchie nasekomoyadnye (Mammalia, Inectivora). Publishing House Nauka, 4, Leningrad. (In Russian) HELLER F. 1958. Eine neue altquartäre Wirbeltierfauna von Erpfingen (Schwäbische Alb). Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, 107: 1-102. HERMANN J. 1780. [In:] E.A.W. ZIMMERMANN 1780. Geographische Geschichte des Menschen, 2, p. 6. Weygandschen Buchhandlung, Leipzig, 344 pp. HINTON M.A.C. 1911. The British fossil shrews. Geological Magazine, 8: 529-539. HÍR J., MÉSZÁROS L. G. 2002. Middle Miocene insectivores and rodents (Mammalia) from Sámsonháza (Northern Hungary). Fragmenta Palaeontologica Hungarica, 20: 9-23. HUTTERER R. 1995. Archaeodesmana TOPACHEVSKY and PASHKOV, the correct name for Dibolia RÜMKE, a genus of fossil water moles (Mammalia: Talpidae). – Bonner Zoologische Beitrage, 45: 171-172. JÁNOSSY D. 1986. Pleistocene vertebrate faunas of Hungary. Akadémiai Kiadó, Budapest, 208 pp. KERR R. 1792. Animal kingdom, or zoological system, of the celebrated Sir Charles LINNAEUS. (Class I. Mammalia: containing a complete systematic description, arrangement, and nomenclature, of all known species and varieties of Mammalia, or animals which suck to their young; being a translation of that part of the Systema Naturae, as lately published, with great improvements, by Professor GMELIN of Goettingen). B. RZEBIK-KOWALSKA, L. I. REKOVETS 168
KORMOS T. 1913. Trois nouvelles espèces fossiles des desmans en Hongrie. Annales Musei Nationales Hun- garici, 11: 135-146. KORMOS T. 1926. Amblycoptus oligodon n. g. & n. sp., eine neue Spitzmaus aus dem Ungarischen Pliozän. An- nales Musei Nationalis Hungarici, 24: 369-391. KORMOS T. 1930. Diagnosen neuer Säugetiere aus der oberpliozänen Fauna des Somlyóberger bei Püspök- fürdö. Annales Musei Nationalis Hungarici, 27: 237-246. KORMOS T. 1934. Neue Insektenfresser, Fledermäuse und Nager aus dem Oberpliozän der Villanyer Gegend. Földtani Közlöny, 64: 296-321. KOWALSKI K. 1956. Insectivores, bats and rodents from the Early Pleistocene bone breccia of Podlesice near Kroczyce (Poland). Acta Palaeontologica Polonica, 1: 331-394. KRETZOI M. 1938. Die Raubtiere von Gombaszög nebst einer Ubersicht der Gesamtfauna. (Ein Beitrag zur Stratigraphie des Altquartaers). Annales Historico-naturales Musei Nationalis Hungarici, 1937-1938, 31: 88-157. LAXMANN E. 1788. Sorex caecutiens. P. 1. Natural History of Central Asia. Nova Acta, Academia of Sciences, Petropoli, 3: 54-83. LINNAEUS C. 1758. Systema naturae pre regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Laurentius Salvus, Stockholm, 824pp. LINNAEUS C. 1766. Systema naturae per regna tria nature, secundum classes, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Editio 12, 1: 1-532. LUNGU A.N. 1981. Gipparionovaya fauna Srednego Sarmata Moldavii (nasekomoyadnye, zaitseobraznye i gryzuny). Publishing House Shtiintsa, Kishinev. (In Russian). MARKOVA A.K. 1980. Rannepleistotsenovaya mikroteriofauna Basse0na Dona i eN sravnenie s melkimi mle- kopitayushchimi Pridneprovya. Publishing House Nauka: 107-139. (In Russian). MARKOVA A.K. 1985. Mikroteriofauna Mikulinskogo mezhlednikovya (po materialam Russko0 Ravniny). – Doklady Akademii Nauk SSSR, 284: 718-721. (In Russian). MARKOVA A.K. 1987. Paleoclimatic reconstruction of the Late Pleistocene in the upper and middle Dnieper regions and in Byelorussia from data on small mammals. [In:] M. PÉSCI, A. A. VELICHKO (eds) – Paleoge- ography and Loess, Akadémiai Kiadó, Budapest: 89-97. MEZHZHERIN V.A. 1972. Zemlero0ki-burozubki (Sorex, Insectivora, Mammalia) iz ple0stotsenovykh otloz- henii SSSR. Teriologiya 1. Publishing House Nauka, Siberian Branch, Novosibirsk. (In Russian). MEZHZHERIN V.O., SVISTUN V.I. 1966. Novii pidvid vikopno0 burozubki zvicha0no0 Sorex araneus praetetra- gonurus subsp. nov. Dopovidi Akademii Nauk URSR, 8: 1071-1074. (In Ukrainian). MEZHZHERIN V.A., SVISTUN V.I. 1968. O rodstvennykh svyazyakh mezhdu iskopaemo0 burozubko0 iz sred- negolotsenovykh otlozhenii r. Desny i retsentno0 burozubko0 arktichesko0. Vestnik Zoologii, 1: 87-89. MILLER G.S. 1901. Five new shrews from Europe. Proceedings of the Biological Society of Washington, 14: 41-45. NESIN V.A. 2013. Neogenovye Murinae (Rodentia, Muridae) Ukrainy. Monograph. Universitetskaya Kniga, Kiev. (In Russian). NESIN V.A., NADACHOWSKI A. 2001. Late Miocene and Pliocene small mammal faunas (Insectivora, Lago- morpha, Rodentia) of Southeastern Europe. Acta zoologica cracoviensia, 44: 107-135. NOVACEK M.J. 1986. The skull of leptictid insectivorans and the higher-level classification of eutherian mam- mals. Bulletin of the American Museum of Natural History, 183: 1-112. PALLAS P.S. 1811 [1831]. Zoographia Rosso-Asiatica, sistens omnium Animalium in extenso Imperio Ros- sico et adjacentibus maribus observatorum recensionem, domicillia, mores et descriptions, anatomem atque icons plurimorum. [ICZN Opinion 212 – dates of volumes: 1 & 2 : 1811, 3: 1814]. Petropoli, in officina Caes. Academiae scientiarum 1. PASHKOV A.V., TOPACHEVSKY V.A. 1990. Novye predstaviteli roda Desmana (Insectivora, Talpidae) iz pozdnepliotsenovykh otlozhenii tsentralno0 Evropy i yugo-zapada evrope0sko0 chasti SSSR. Vestnik Zoolo- gii, 5: 25-34. (In Russian). PENNANT T. 1771. Synopsis of Quadrupeds. J. MONK, Chester, 382 pp. PETÉNYI S. J. 1864. Hátrahagyott Munkái. F. Eggenberger, Budapest, 130 pp. PIDOPLICHKO I.G. 1955. Novye dannye o faune pozvonochnykh antropogenovykh otlozhenii Ternopolsko0 oblasti. Doklady Akademii Nauk SSSR, 100: 989-991. (In Russian). Recapitulation Ukrainian fossil insectivores 169
PIDOPLICHKO I.G. 1956. Do vivchennya fauni antropogenovikh khrebetnikh Ternopilsko0 oblasti. Naukovi zapiski Prirodoznavchogo Muzeyu Lvivskogo Filialu AN URSR, 5: 45-53. (In Ukrainian). PRIETO J., VAN DEN HOEK OSTENDE L., HÍR J. 2012. The Middle Miocene insectivores from Sámsonháza 3 (Hungary, Nógrád County): Biostratigraphical and palaeoenvironmental notes near to the Middle Miocene Cooling. Bulletin of Geosciences, 87: 227-240. QIU Z.D., STORCH G. 2000. The early Pliocene Micromammalian Fauna of Bilike, Inner Mongolia, (Mam- malia: Lipotyphla, Chiroptera, Rodentia, Lagomorpha). Senckenbergiana lethaea, 80(1): 173-229. QIU Z.D., STORCH G. 2005. China. [In:] L. W. VAN DEN HOEK OSTENDE, C. S. DOUKAS, J. W. F. REUMER (eds) – The fossil record of the Eurasian Neogene insectivores (Erinaceomorpha, Soricomorpha, Mam- malia), Part. I. Scripta Geologica, Special Issue, 5: 37-50. RABEDER G. 1972. Die Insectivoren und Chiropteren (Mammalia) aus dem Altpleistozän von Hundsheim (Niederösterreich). Annalen des Naturhistorischen Museums, 76: 375-474. REKOVETS L.I. 1990. La localité Bolchévik 2 et l’évolution de la microthériofaune au PléistocPne dans la rJgion Septentrionale de la Mer Noir. L’Anthropologie, 94: 659-674. REKOVETS L.I., CHEPALYGA A.L., NESIN V.A., SVETLITSKAYA T.V. 1990. Bolshevik 2 – novoe mesto- nakhozhdenie mlekopitayushchikh antropogena Severnogo Prichernomorya. [In:] I. A. AKIMOV (ed.) – No- vosti faunistiki i sistematiki, Naukova Dumka, Kiev: 175-180. (In Russian). REKOVETS L.I., NADACHOWSKI A. 1995. Pleistocene voles (Arvicolidae) of the Ukraine. Paleontologia i Evolutiò: 145-245. REKOVETS L.I., PASHKOV A.V. 2009. Novye mestonakhozhdeniya mikroteriofauny pozdnego neogena Ukrainy. Zbirnik Naukovikh Prats Institutu Geologichnikh Nauk NAN Ukraini, Kiev: 355-360. (In Russian). REKOVETS L.I., STARKIN A.V. 1990. Teriofauna pozdnepaleoliticheskogo poseleniya Anetovka-2 yuga Ukrainy. Vestnik Zoologii, 3: 40-44. (In Russian). REUMER J.W.F. 1984. Ruscinian and Early Pleistocene Soricidae (Insectivora, Mammalia) from Tegelen (The Netherlands) and Hungary. Scripta Geologica, 73: 1-173. RÜMKE C.A. 1976. Insectivora from Pikremi and Biodrak (Greece). Proceedings of the Koninklijke Neder- landse Akademie van Wetenschappen, 79: 256-270. RÜMKE C.A. 1985. A review of fossil and recent Desmaninae (Talpidae, Insectivora). Utrecht micropaleonto- logical bulletins. Special publication, 4, Utrecht. RZEBIK-KOWALSKA B. 1975. The Pliocene and Pleistocene insectivores (Mammalia) of Poland. II. Soricidae: Paranourosorex and Amblycoptus. Acta zoologica cracoviensia, 20: 167-182. RZEBIK-KOWALSKA B. 1976. The Neogene and Pleistocene insectivores (Mammalia) of Poland. III. Sorici- dae: Beremendia and Blarinoides. Acta zoologica cracoviensia 22: 359-385. RZEBIK-KOWALSKA B. 1981. The Pliocene and Pleistocene Insectivora (Mammalia) of Poland. V. Soricidae: Neomysorex n. g. and Episoriculus ELLERMAN et MORRISON-SCOTT, 1951. Acta zoologica cracoviensia, 25: 227-250. RZEBIK-KOWALSKA B. 1990. Pliocene and Pleistocene Insectivora (Mammalia) of Poland. VI. Soricidae: Deinsdorfia HELLER, 1963 and Zelceina SULIMSKI, 1962. Acta zoologica cracoviensia, 33: 45-77. RZEBIK-KOWALSKA B. 1991. Pliocene and Pleistocene Insectivora (Mammalia) of Poland. VIII. Soricidae: Sorex LINNAEUS, 1758, Neomys KAUP, 1829, Macroneomys FEJFAR, 1966, Paenelimnoecus BAUDELOT, 1972 and Soricidae indeterminata. Acta zoologica cracoviensia, 34: 323-424. RZEBIK-KOWALSKA B. 1996. Insectivora (Mammalia) from the Miocene of Be³chatów in Poland. III. Dimyli- dae SCHLOSSER, 1887. Acta zoologica cracoviensia 39: 447-468. RZEBIK-KOWALSKA B. 1998. Fossil history of shrews in Europe. [In:] J. M. WÓJCIK, M. WOLSAN (eds) – Evolution of shrews. Mammal Research Institute, Polish Academy of Sciences, Bia³owie¿a: 23-92. RZEBIK-KOWALSKA B. 2009. Biodiversity of Polish fossil insectivores (Erinaceomorpha, Soricomorpha, In- sectivora, Mammalia) compared to the European and global faunas. Institute of Systematics and Evolution of Animals Polish Academy of Sciences, Kraków, 123 pp. RZEBIK-KOWALSKA B., NESIN V. A. 2010. Erinaceomorpha and Soricomorpha (Insectivora, Mammalia) from the Late Miocene of Ukraine. Institute of Systematics and Evolution of Animals Polish Academy of Sciences, Kraków, 61 pp. RZEBIK-KOWALSKA B., PAW£OWSKI, J. 1994. Ruemkelia (Mammalia, Insectivora, Talpidae) nom. nov. from Dibolia RÜMKE, 1985 (nec LATREILLE, 1829). Acta zoologica cracoviensia, 37: 75-76. RZEBIK-KOWALSKA B., REKOVETS L. I. (in press). New data on Eulipotyphla (Insectivora, Mammalia) from the Late Miocene to the Middle Pleistocene of Ukraine. Palaeontologica Electronica. B. RZEBIK-KOWALSKA, L. I. REKOVETS 170
RZEBIK-KOWALSKA B., TOPACHEVSKY V. A. 1997. Insectivora (Mammalia) from the Miocene of Grytsiv in Ukraine. I. Heterosoricidae VIRET and ZAPFE, 1951. Acta zoologica cracoviensia, 40: 27-247. SCHLOSSER M. 1887. Die Afen, Lemuren, Chiropteren, Insectivoren, Creodonten und Carnivoren des eu- ropäischen Tertiärs und deren Besiehungen zu ihren lebeden und fossilen aussereuropäischen Verwandten. Beiträgr zur Paläontologie Österreich-Ungarns und des Orients 6 (1888). SCHMIDT-KITTLER N. 1973. Dimyloides – Neufunde aus der oberoligozänen Spaltufüllung „Ehrenstein 4“ (Süddeutschland) und die Systematische Stellung der Dimyliden (Insectivora, Mammalia). Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie, 13: 115-139. SCHREUDER A. 1940. A revision of the fossil water-moles (Desmaninae). Archives Néerlandaises de Zoolo- gie, 4: 201-333. SHEVCHENKO A.I. 1965. Opornye kompleksy melkikh mlekopitayushchikh pliotsena i nizhnego antropogena yugo-zapadno0 chasti Russko0 Ravniny. Stratigraficheskoe znachenie antropogenovo0 fauny melkikh mle- kopitayushchikh. Publishing House Nauka: 7-58. (In Russian). SICKENBERG O. 1971. Revision der Wirbeltierfauna der Höhle Petralona (Griech, Mazedonien). Annales Géologique des Pays Helléniques, 23: 230-264. SOMMER R.S. 2007. When east met west: the sub-fossil footprints of the west European hedgehog and the northern white-breasted hedgehog during the Late Quaternary in Europe. Journal of Zoology, 273: 82-89. STORCH G. 1978. Die turolische Wirbeltierfauna von Dorn-Dürkheim, Rheinhessen (SW-Deutschland). 2. Mammalia: Insectivora. Senckenbergiana Lethaea, 58: 421-449. STORCH G., QIU Zh., ZAZHIGIN V.S. 1998. Fossil history of shrews in Asia. [In:] J. M. WÓJCIK, M. WOLSAN (eds) – Evolution of shrews. Mammal Research Institute, Polish Academy of Sciences, Bia³owie¿a: 93-120. STORCH G., ZAZHIGIN V.S. 1996. Taxonomy and phylogeny of the Paranourosorex lineage. Neogene of Eurasia (Mammalia: Soricidae: Anourosoricini). Paläontologische Zeitschrift, 70: 257-268. SVISTUN V.I. 1968. Fauna pozdneantropogenovykh mlekopitayushchikh Romankovskogo allyuvialnogo me- stonakhozhdeniya. [In:] I. G. PIDOPLICHKO (ed.) – Prirodnaya obstanovka i fauny proshlogo. Publishing House Naukova Dumka, Kiev: 3-56. (In Russian).0 TATARINOV K.A. 1958. Znakhidki zemleri0ki blyarini na Ukraini. Dopovidi Akademii Nauk URSR, 1: 81-84. (In Ukrainian). TATARINOV K.A. 1965a. Ple0stotsenovye mlekopitayushchie iz Nizhnekrivchansko0 peshchery (Podoliya). Paleontologicheskii Sbornik Lvovskogo Gosudarstvennogo Universiteta im. I. FRANKO, 2: 30-37. (In Russian). TATARINOV K.A. 1965b. Neogenovye pozvonochnye zapadnykh oblaste0 Ukrainy. Doklady i Soobshcheniya Lvovskogo otdela geograficheskogo obshchestva USSR, Lvov: 18-21. ( In Russian). TATARINOV K.A. 1970. Fauna neogenovykh i antropogenovykh pozvonochnykh Podolii i Prikarpatya, eN isto- riya i sovremennoe sostoyanie. Unpublished PhD Thesis, Akademiya Nauk Ukrainsko0 SSR, Kiev. (In Rus- sian). TOPACHEVSKY V.O. 1956. Reshtki dribnogo verblyuda (Paracamelus alutensis) z verkhnopliotsenovikh vik- ladiv Pivdnya URSR. Trudi Institutu Zoologii ANU RSR, 13: 93-100. (In Ukrainian). TOPACHEVSKY V.O. 1957a. Piznople0stotsenova ta golotsenova fauna ssavtsiv z suchasnikh alyuvialnikh vidkladiv nizhnogo Dnipra. Trudi Institutu Zoologii ANU RSR, 14: 113-128. (In Ukrainian). TOPACHEVSKY V.O. 1957b. Do vivchennya fauni piznopliotsenovikh ta rannoantropogenovikh khrebetnikh z davnikh alyuvialhikh vidkladiv pivdnya URSR. Trudi Institutu Zoologii ANU RSR, 14: 141-147. (In Ukrainian). TOPACHEVSKY, V.O. 1959. Nova forma vykhukhola z serednogo ta piznogo ple0stotsenu URSR. Dopovidi Akademii Nauk Ukrainsko0 RSR, 6: 666-671. (In Ukrainian). TOPACHEVSKY V.A. 1961a. Novy0 pliotsenovy0 vid vykhukholi iz Predkavkazya. Paleontologicheskii Zhur- nal, 4: 131-137. (In Russian). TOPACHEVSKY V.O. 1961b. Novi materiali do antropogenovo0 istorii fauni ssavtsiv Serednogo Pridniprovya. Zbirnik Prats Zoologichnogo Muzeyu, Kiev, 30: 9-20. (In Ukrainian). TOPACHEVSKY V.O. 1962a. Vikopni vikhukholi rodu Desmana z neogenovikh ta antropogenovikh vidkladiv evropeisko0 chastini SRSR. [In:] I. G. PIDOPLICHKO (ed) - Vikopni fauni Ukraini i sumizhnikh teritorii, Akademiya Nauk Ukrainsko0 RSR, Kiev: 5-90. (In Ukrainian). TOPACHEVSKY V.O. 1962b. Do pitannya pro geologichnii vik davnikh alyuvialnikh vidkladiv pivnichno0 dily- anki nizhnogo Dnipra. Geologichnii Zhurnal, 22: 106-109. (In Ukrainian). TOPACHEVSKY V.A. 1965. Nasekomoyadnye i gryzuny noga0sko0 pozdnepliotsenovo0 fauny. Publishing House Naukova Dumka, Kiev. (In Russian). Recapitulation Ukrainian fossil insectivores 171
TOPACHEVSKY V.O. 1966. Novii rid krupno0 bilozubo0 zemleri0ki (Insectivora, Soricidae) z pliotsenovikh vidkladiv pivdnya Ukraini. [In:] Ekologia ta istoria khrebetnych fauny Ukrainy, Publishing House Naukova Dumka, Kiev: 90-96. (In Ukrainian). TOPACHEVSKY V.O., CHEPALIGA A.L., NESIN V.A., REKOVETS L.I., TOPACHEVSKY I.V. 1988. Mikroterio- fauna (Insectivora, Lagomorpha, Rodentia) lektostratotipu Pontu. Dopovidi Akademii Nauk Ukrainskoi RSR, 4: 73-76. (In Ukrainian). TOPACHEVSKY V.A., KORNIETS N.L., SVISTUN V.I. 1975. Novye dannye k paleoteriologichesko0 kharakter- istike Tiraspolskogo (Mindelskogo) yarusa v sostave Khadzhibe0sko0 svity. Vestnik Zoologii, 1: 67-73. (In Russian). TOPACHEVSKY V.A., NESIN V.A., PRISYAZHNYUK V.A. 1990. Svoeobraznoe soobshchestvo melkikh mle- kopitayushchikh perekhodno0 k Pontu zony Meotisa yugo-zapada USSR. Doklady Akademii Nauk Ukrain- sko0 SSR, 9: 77-80. (In Russian). TOPACHEVSKY V.A., NESIN V.A.,, TOPACHEVSKY I.V. 1997. Ocherk istorii mikroteriofaun (Insectivora, La- gomorpha, Rodentia) Ukrainy v otrezke vremeni srednii Sarmat – Akchagyl. Vestnik Zoologii, 31: 3-14. (In Russian). TOPACHEVSKY V.A., PASHKOV A.V. 1983. Nadvidovaya sistematika vykhukhole0 roda Desmana (Insecti- vora, Talpidae), Vestnik Zoologii, 3: 39-45. TOPACHEVSKY V.A., PASHKOV A.V. 1990. Novye predstaviteli roda Desmana (Insectivora, Talpidae) iz eo- ple0stotsenovykh otlozhenii yuga evroope0sko0 chasti SSSR. Vestnik Zoologii, 1: 28-38. (In Russian). TOPACHEVSKY V.A., SKORIK A.F., CHEPALYGA A.A. 1979. Novye materialy po faune mollyuskov i melkikh mlekopitayushchikh pozdnego pliotsena Odesskogo Kuyalnika. Vestnik Zoologii, 5: 11-18. (In Russian). VAN DEN HOEK OSTENDE L.W. 1995. Insectivore faunas from the Lower Miocene of Anatolia. Part 3: Dimyli- dae. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, 98: 19-38. VAN DEN HOEK OSTENDE L.W., FURIÓ M. 2005. Spain. – [In:] L.W. van den HOEK OSTENDE, C.S. DOU- KASAND, J.W.F. REUMER (eds) – The fossil record of the Eurasian Neogene insectivores (Erinaceomorpha, Soricomorpha, Mammalia). Part. I. Scripta Geologica Special Issue 5: 148-284. VIRET J., ZAPFE H. 1951. Sur quelques Soricidés miocènes. Eclogae Geologicae Helvetiae, 44: 411-426. VON KOENIGSWALD W. 1970. Mittelpleistozäne Kleinsäugerfauna aus der Spaltenfüllung Petersbuch bei Eichstätt. Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie, 10: 407-432. WADDELL P.J., OKADA N., HASEGAWA M. 1999. Towards resolving the interordinal relationships of placen- tal mammals. Systematic Biology, 48: 1-5. ZAITSEV M.V., BARYSHNIKOV G.F. 2002. Pleistocene Soricidae (Lipotyphla, Insectivora, Mammalia) from Treugolnaya Cave, Northern Caucasus, Russia. Acta zoologica cracoviensia, 45: 283-305. ZIEGLER R. 1999. Order Insectivora. [In:] E. ROOSNER, K. HEISSIG (eds) – The Miocene land mammals of Europe, Verlag Dr Friedrich Pfeil, Munchen: 53-74.